One female specimen of the cetacean harbour porpoise Phocoena Phocoena (Linnaeus, 1758) was found dead on the North Sea coast of Denmark (Jutland) in May 2025. During necropsy severe pathological changes of the liver were observed (hepatic fibrosis, enlarged and thickened biliary duct walls) and when dissecting the biliary ducts six intact trematodes (total length 6.9-7.9 mm, width 1.9-2.0 mm) were recovered. Following conservation in formalin four parasites were haematoxylin stained, one specimen kept unstained but all five mounted for morphological analysis. The sixth specimen conserved in 70% ethanol was subjected to DNA purification and subsequent PCR and sequencing. All the parasites were identified as fully adult Campula oblonga (Cobbold, Trans Linnean Soc Lond 22(3):155-172, 1858). Morphometric, morphological and molecular data (NADH dehydrogenase, subunit 3 (ND3), mtDNA) are presented and indicate close relation with previous isolates from the North and Baltic Seas but a lower similarity to Pacific conspecifics.
Accurate species-level classification of prokaryotic 16S rRNA sequences remains difficult: existing tools rely on exact alignment, k-mer heuristics, or phylogenetic placement and are limited by incomplete reference databases. Deep learning approaches in microbial genomics have focused largely on whole-genome metagenomics, leaving 16S taxonomy under-supported. We present DeepTaxa, a hybrid CNN-BERT framework that pairs a multiscale CNN with a transformer trained from scratch on the DNABERT-2 BPE vocabulary, producing parallel rank-specific predictions across the seven Linnean ranks. On the Greengenes2 2024.09 test set, DeepTaxa achieves species-level accuracy of 92.96% and F1 of 0.9212 (3-seed mean; cross-seed standard deviation ≤ 0.0008 F1 at every rank), with F1 above 0.99 from domain through class and a species-level expected calibration error of 0.0242. DeepTaxa exceeds DADA2 (90.05%) and QIIME 2 (85.01%) at the species rank on the same held-out test set, with larger gains over the k-mer-based classifiers SINTAX and Kraken 2. Performance degrades smoothly with decreasing training-set similarity (species F1 from 0.95 to 0.45), and a dedicated V3-V4 amplicon checkpoint reaches 87.55% species accuracy from an approximately 420 bp window. Source code, trained checkpoints for full-length 16S and V3-V4 amplicons, curated datasets, and reproducible workflows are publicly available at github.com/systems-genomics-lab/deeptaxa and huggingface.co/systems-genomics-lab/deeptaxa.
DNA barcode reference libraries provide useful tools for specimen identification, highlighting potential new species and detecting introduced ones. Here, we present a comprehensive DNA barcode library for European ants and, in order to tackle the Linnean, Wallacean and Darwinian shortfalls of this group, we provide an updated checklist, distribution data, mitochondrial genetic diversity maps and mitochondrial gene trees. The European ant fauna is here established to include 55 genera and 650 species (587 of which are native), including one species newly recorded for Europe and novel citations for 26 species from 11 countries. Our genetic dataset includes 6530 georeferenced COI sequences (62.1% de novo) for 506 species (77.8%) across all genera. On average, 12.9 sequences were obtained per species, and 209 species were sequenced for the first time. We generated intra- and interspecific genetic distance estimates, 52 genus-level trees, mitochondrial genetic diversity and specimen maps for 384 species, as well as haplotype networks for 289 species, available in the Atlas V1.0 'The Mitochondrial Genetic Diversity Maps of European Ants'. We estimate that 56.3% of European ants are monophyletic with respect to the COI gene and can be unambiguously identified by DNA barcoding, though performance varies widely among genera. We observed moderate levels of barcode sharing (19.3%) and of barcode gap presence (47.6%), as well as high levels of intraspecific divergences (up to 17.9%). These findings likely reflect both biological and operational factors and highlight the existence of potential cryptic taxa and the need for taxonomic revisions. The framework presented here aims to facilitate future research, species discovery and conservation of European ants.
BACKGROUND: Value-Based Healthcare (VBHC) emphasises delivering high-quality care measured by patient outcomes rather than the volume of services. Although mostly partially implemented in civilian health systems, VBHC integration within military health systems (MHS) lags behind. Given current geopolitical developments and the unique operational demands of military healthcare, this study aimed to assess the desirability and feasibility of implementing VBHC in the Dutch MHS. METHODS: A cross-sectional, descriptive survey was conducted in 2024 among active-duty personnel in the Netherlands Armed Forces. Stratified sampling targeted three stakeholder groups—care recipients, care providers, and care facilitators—followed by voluntary response recruitment. A self-administered online questionnaire assessed familiarity with VBHC, perceived desirability and feasibility of its implementation, and prioritised components for application. The survey was structured around case-based scenarios derived from the Linnean VBHC framework, adapted to the military context. Quantitative data were analysed using descriptive statistics and Likert-scale medians; open-ended responses were analysed thematically. RESULTS: Of 912 eligible participants, 290 completed the survey (32% response rate), comprised of 45% care recipients, 33% care providers, and 22% care facilitators. Most respondents (67%) had over 15 years of military service; 58% held higher education degrees. Familiarity with VBHC increased throughout the survey. Overall, 96% of respondents perceived VBHC as valuable for the Dutch MHS. Notably, 64% believed VBHC could benefit both regular and operational care. A collaborative implementation approach involving all three stakeholder groups was preferred by 82%. However, concerns were raised about feasibility in large-scale combat operations, highlighting the need to align VBHC components with military readiness and mission-critical objectives. CONCLUSION: This study provides initial evidence supporting broad acceptance of VBHC within the Dutch MHS, particularly for non-operational military healthcare contexts. While desirability was high, practical applicability raised concerns, especially regarding combat and operational care settings. These findings suggest the potential for VBHC integration, but emphasize the necessity for a military-specific adaptation. Future research should explore targeted implementation strategies that balance patient-centered outcomes with the unique operational demands of military healthcare environments.
The names published by Linnaeus in Species Plantarum represent the foundation of modern plant taxonomy. Despite their systematic value, very few of Linnaeus's original type specimens have been analyzed using current DNA sequencing technologies. Here, we performed high-throughput sequencing on Linnean and other type specimens of Ulva, a genus of ecological and commercial importance. Chloroplast and mitochondrial genomes were assembled for Linnaeus's U. compressa, U. intestinalis, U. lanceolata, and U. linza as well as Kützing's Phycoseris smaragdina type specimens. Phylogenetic analyses of these data showed that the names U. compressa and U. intestinalis were correctly applied, but U. linza and U. lanceolata were misapplied. Ulva linza is the earliest available name for the European species currently called U. pseudocurvata. The correct name for the globally distributed species previously known as U. "linza" is Ulva smaragdina (Kützing) comb. nov. The names Ulva lanceolata, U. crispata, and Phycoseris olivacea do not represent distinct species, instead being heterotypic synonyms of U. compressa, and P. planifolia is a heterotypic synonym of U. intestinalis. These results demonstrate that genetic characterization of type material can unequivocally resolve longstanding taxonomic debates over scientific names.
Research Highlight: Colares, L. F., Peres, C. A., Dambros, C. S. (2026). Life history induces markedly divergent insect responses to habitat loss. Journal of Animal Ecology. https://doi.org/10.1111/1365-2656.70117. Habitat loss is driving biodiversity collapse worldwide. Although this phenomenon has been extensively studied across many taxa and regions, we still lack information about whether species with distinct life histories respond differently to habitat loss. This challenge is particularly critical for tropical insects, where knowledge gaps remain large due to the Linnean (taxonomy) and Raunkiæran (traits) shortfalls. In this issue, Colares et al. (2025) address these gaps by using 236 sticky traps across the world's largest man-made tropical forest archipelago in the Central Amazon (~360,000 ha), generating a dataset of ~23,000 individual insects. They combined these surveys of insect fauna with computer vision models to assess how habitat loss affects both α- and β-diversity in insects with contrasting life histories (terrestrial vs. aquatic). The study reveals that responses diverge strongly depending on whether taxa rely on terrestrial or aquatic environments during their ontogeny. Whereas low forest amount reduced the number of terrestrial species, it increased species with aquatic life histories. Importantly, the authors also linked insect responses to body size (a proxy for dispersal ability), suggesting that larger insects, which disperse more successfully across the water matrix, may be favoured as 'winner' species in fragmented habitats. The findings of Colares et al. (2025) have broad implications for animal ecology and insect conservation. First, they highlight that insect declines in response to habitat loss are largely driven by traits that confer high or low resilience to reductions in forest cover. Second, they underscore the potential of computer vision as a powerful tool for uncovering key information about insect populations, thereby facilitating applied research such as rapid biodiversity surveys and long-term monitoring.
Very low-intensity selective logging can be a compromise between strict conservation and income-generating land use in tropical forests. Investigating how selective logging influences the understory environment and seedling dynamics as the forest regenerates offers insights into whether logging alters forest dynamics, influencing the composition and structure of future forests. We explored how very low-intensity logging (<2 trees ha-1) influences understory factors and seedling dynamics across a logging chronosequence (unlogged forest vs. actively logged forest and forest logged 4 and 14 years prior). To do this, we assessed (1) how canopy openness, prevalence of vegetation damage, and elephant trails differ in logged forests at different recovery stages compared to unlogged forest; (2) how these understory factors influence seedling dynamics; (3) how seedling dynamics differ across the logging chronosequence; and (4) how logging impacts liana vs. tree seedlings across the chronosequence. We observed greater canopy openness and vegetation damage in logged forests up to 4 years after logging and higher elephant trail prevalence 14 years after logging compared to unlogged forests. Seedling survival was lower in plots with higher canopy openness, more vegetation damage, and on elephant trails, while growth and recruitment were not affected by these variables. Actively logged forests initially had lower seedling survival and recruitment, but higher growth rates compared to unlogged forests. However, 14 years after logging, seedling dynamics were mostly similar to unlogged forests. Liana seedlings had a slight growth advantage over tree seedlings in all logged forests compared to unlogged forests. Results from our study suggest that very low-intensity selective logging causes temporary shifts in understory dynamics rather than long-term shifts in forest recovery trajectories. These managed areas have potential as land that can contribute to OECM targets-functioning as mixed-use corridors, connecting protected areas across a landscape and contributing to biodiversity and wildlife conservation-especially in countries with high forest cover and low deforestation.
Trehalose 6-phosphate (T6P), a trehalose synthesis intermediate and sugar phosphate, serves as a signaling molecule coordinating sucrose status with plant growth and development. Beyond its metabolic role, the T6P pathway integrates exogenous and other endogenous cues to regulate key developmental transitions, including embryogenesis, seed maturation and filling, shoot branching, vegetative and reproductive phase transitions, and tuber and lateral root formation. Dynamic spatiotemporal expression patterns of T6P-pathway genes correlate with developmental stages, though their specific contributions to the initiation and progression of these transitions remain under investigation. Here, we provide recent insights and future perspectives on the T6P pathway, emphasizing its role in orchestrating diverse plant developmental programs across model and crop species and highlighting emerging mechanistic insights into its functions.
CYSTEINE-RICH RECEPTOR-LIKE PROTEIN KINASEs (CRKs) play an important role in plant development and stress responses. One of the best described members of the Arabidopsis CRK family is CRK2, which was proposed as a crucial regulator of intercellular transport facilitated by plasmodesmata (PD). As intercellular channels allowing symplastic communication, PD-mediated transport is predominantly regulated by callose synthase (CALS)-mediated callose deposition. This process can impact not just the distribution of molecules between adjacent cells, but also the symplastic loading of vascular tissue, thereby influencing plant stress responses and developmental processes. Here we described the overlapping expression pattern of genes encoding phylogenetically closely related CALS1 and CALS3. Both CALSs were phosphorylated in vitro by CRK2, and the genetic interaction between genes encoding CRK2 and CALS1 or CALS3 revealed their impact on callose deposition, rosette growth, primary root length, and development, represented as a decreased number of true leaves. Importantly, we observed significant accumulation of starch in crk2 mutant plants, especially in developmentally older leaves, which was reverted by the independent introduction of cals1.5 and cals3.1 into the crk2 mutant background. The observed starch accumulation was accompanied by photosynthesis inhibition. We propose that the growth and developmental alterations of crk2 are caused by decreased phloem loading, which resulted in starch accumulation in source organs, and subsequent sink tissue starvation. Our results propose CRK2 as negative regulator of CALS1 and CALS3 regulating source to sink transport, which impacts plant growth and development.
Plants continuously emit volatile organic compounds (VOCs), which can influence the physiology and behavior of neighboring plants. While the ecological role of stress-induced VOCs is well established, the function of constitutive VOCs released by undamaged plants in mediating plant-plant interactions remains less understood. Here, we demonstrate that barley plants can detect the growth rate of undamaged conspecific neighbors through constitutive VOCs and respond by modulating their growth-defense trade-off accordingly. Exposure to volatiles from cultivars with contrasting growth (slow or fast) triggered distinct shifts in biomass accumulation and gene expression in receiver plants, whereas VOCs from cultivars with similar growth rates had negligible effects. Transcriptomic analysis revealed cultivar-specific transcriptional reprogramming of growth- and defense-related pathways, suggesting that constitutive VOCs convey information about emitter identity and competitive vigour that receiver plants use to adaptively reallocate resources and prime stress responses in anticipation of competition. These findings uncover a previously unrecognized role of constitutive VOCs as reliable cues of emitter identity and vigor, mediating adaptive responses in neighboring plants under competitive scenarios.
Grindelia mutabilis (Asteraceae, Astereae), a new species from Brazil endemic to the Espinal Ecoregion of the Río de La Plata Grasslands Bioregion and Pampa Province of the Chaco Biogeographical Domain, is proposed and illustrated. The new species is characterized by a combination of traits: small, rosette cespitose habit, linear to linear-oblanceolate leaves, light-yellow to pastel salmon ray florets, three-winged ray floret cypselae bearing a pappus of two to four elements and two-winged disc floret cypselae bearing a pappus of two elements. It has a highly restricted habitat and is known exclusively within Parque Estadual do Espinilho in Rio Grande do Sul, Brazil. Preliminary conservation assessments classify the new species as Critically Endangered. We provide illustrations and photographs, as well as a distribution map with an identification key for the South American Grindelia species with winged cypselae. The intriguing morphology of this species combines characters traditionally regarded as diagnostic for Notopappus, a genus segregated from Haplopappus and Grindelia. Previously published phylogenetic studies of related taxa indicate that the recognition of Notopappus as monophyletic is not supported and render Grindelia as non-monophyletic too. Based on this combined morphological evidence and existing phylogenetic hypotheses, we reaffirm the non-monophyly of Notopappus and formally propose its synonymization under Grindelia s.l.
The earliest available names for sponges were proposed by Linnaeus in volume II of the 10th edition of his Systema Naturae published in 1759. He described the eleven Spongia species cursorily, and for the illustrations and type localities he referred to pre-1758 works of himself and contemporary authors. Five of the species he erected have been typified by the designation of a lectotype (one species, Spongia lacustris) and neotypes (four species, Spongia officinalis, S. fistulosa, S. muricata and S. fluviatilis), but the remaining six species lacked extant type material and no neotypes have been designated so far (S. oculata, S. nodosa, S. flabelliformis, S. tubulosa, S. aculeata, and S. infundibuliformis). It is the purpose of this study to review and further typify the 1759 Linnean species. In order to clarify the morphological and nomenclatural issues, I traced the history of all eleven species in the pre-1758 and early post-1758 literature and evaluated the information upon which Linnaeus based his 1759 species descriptions. Of all species, images cited by Linnaeus and contemporary authors are reproduced and discussed. Photographs of the (neo-)type specimens and reliably identified in situ or preserved non-type specimens are included. For each species, I include a diagnosis, derived from representative descriptions in the recent literature. Distribution maps downloaded from the World Porifera Database provide the occurrence of the species. Where possible, molecular resources available for these first Linnean species are listed. With the six neotypes proposed for the above-mentioned species the typification of all eleven species erected by Linnaeus in 1759 is completed. The typification process of these earliest species is judged to be of importance for the classification of several speciose poriferan clades, Spongia and Spongiidae, Haliclona and Chalinidae, Ianthella and Ianthellidae, Spongilla and Spongillidae, as well as the ubiquitous genera Amphimedon and Ephydatia.
Polyploidization has occurred throughout the tree of life and is particularly common in plants. Despite its ubiquity, our understanding of the short- and long-term effects and consequences of genome doubling in natural populations remains incomplete. In this study, we identified a novel ploidy-variable species system within the ornamental and industrial oilseed genus Orychophragmus (Brassicaceae), which comprises six species, including diploid and tetraploid cytotypes of Orychophragmus taibaiensis. By integrating population-scale genomic and transcriptomic datasets across the species in this genus, we constructed a robust phylogenetic framework and investigated the divergence and demographic history of O. taibaiensis in comparison to its relatives. Specifically, we characterized the geographical distribution patterns of diploids and tetraploids in natural populations of O. taibaiensis, confirmed the autopolyploid origin of tetraploids, and inferred their origin time relative to diploid counterparts. Our findings further revealed that, following genome doubling, tetraploids accumulated a higher genetic load of deleterious mutations, likely due to relaxed purifying selection facilitated by allelic redundancy. Additionally, genome doubling was associated with pronounced changes in gene expression patterns, with differentially expressed genes evolving under relaxed selective constraints. These results highlight that the initial masking of deleterious mutations, changes in expression regulation, and divergent efficacy of selection likely all contribute to shaping the establishment and evolutionary potential of polyploids.
The eusocial honey bee is a model for insect microbiome research, with socially transmitted gut communities that play key roles in health and development. In contrast, solitary bees lack social transmission pathways and often have an environmentally acquired microbiome, which may or may not be functionally important. Using 16S rRNA gene metabarcoding, previous work has described the bacterial communities in the solitary resin bee Megachile tosticauda pollen provisions, brood, and adult bees, but their functional significance has yet to be studied. Here, we investigate the importance of live bacteria for larval development and survival with an antibiotic-feeding experiment, and test whether bacteria are present in adult guts using scanning electron microscopy. Removing live bacteria in the food and gut of feeding larvae had no significant effect on survival or growth. Microscopy revealed no bacterial colonization of the adult gut, and the dominant taxon detected in larval pollen could not be cultured under targeted conditions. These results suggest an absence of a beneficial gut microbiome in M. tosticauda. Based on our findings, we propose that some bacteria detected by DNA-based methods in M. tosticauda may represent relic DNA, surface-associated or transient bacteria. Our findings highlight fundamental differences in the bacterial associations between social and solitary bees.
BACKGROUND: Conserving genetic diversity is crucial for effective germplasm use and crop improvement. Developing core collections with minimal redundancy and maximum diversity requires a clear understanding of population structure. However, the nationwide population structure of moso bamboo (Phyllostachys edulis) remains poorly characterized, creating a major gap for developing representative, non-redundant core collections. RESULTS: Using whole-genome resequencing data from 432 moso bamboo accessions covering a broad geographic range across the distribution of the species in China, we investigated the population genetic structure and diversity patterns. Principal component analysis and phylogenetic tree analyses identified three distinct genetic clusters together with a hybrid group. To identify the optimal strategy for core collection development, we evaluated two stratification schemes, seven sampling strategies, and five sampling intensities. Across 70 candidate cores, stratified sampling combined with expected heterozygosity optimization at 20% intensity (S-HE20) maximized genetic diversity (He = 0.3665; PIC = 0.2904; I = 0.5302) and captured broad phenotypic variation (CR = 82.32%; MD = 0%), yielding an 84-accession core spanning 15 geographic regions. CONCLUSIONS: This study revealed the population genetic structure of moso bamboo and identified the S-HE20 strategy as optimal for core collection construction. The resulting core collection offers a representative and genetically diverse resource for future gene discovery and molecular breeding efforts in moso bamboo.
Plant parasitism is a widespread lifestyle found throughout the plant kingdom that plays important roles in ecology and agriculture. Parasitic plants rely on the formation of specialized parasitic organs called haustoria to invade their hosts and withdraw nutrients. Currently, our knowledge is growing regarding how parasitic plants use haustoria to infect their hosts, modify their physiology and regulate infection. Important factors in plant development are hormone signaling molecules that play essential roles in how plants grow and interact with their surroundings. In recent years, major progress has been made in understanding the relevance of various hormones in plant parasitism. Here, we review recent findings in the field, focusing on the role of hormones in several stages of parasitism, including haustoria induction, vascular development, and interaction with the host. We discuss and compare how hormones influence haustoria development in different parasitic plant lifestyles and species, and identify knowledge gaps in the field. Future work on understanding how hormones influence parasitism is crucial to develop novel ways to control the damage caused by parasitic plants to agriculture, and to discover how parasitic plants efficiently connect to their hosts.
MADS-box transcription factors are a key gene family central to understanding the evolutionary success of land plants. Land plant MADS-box genes are divided into Type I and Type II. They were initially considered orthologous to SRF and MEF2 genes in animals and fungi, respectively, which originated by an ancient duplication before the diversification of extant eukaryotes. However, using updated phylogenetic analyses and AI-based protein structure prediction, we showed that both Type I and Type II genes are plant-specific duplicates derived from MEF2-type ancestors, while ancestral SRF genes were lost before the divergence of the plant lineage. A recent study proposed instead a polyphyletic origin for plant Type I genes. While this study also found that the majority of Type I MADS-box genes in land plants cluster with MEF2 genes, a few genes were grouped as Type I and considered related to SRF genes. By reanalyzing the original data from that study, we do not find supporting evidence for this hypothesis. Moreover, using phylogeny, sequence similarity, and structural evidence, we demonstrate that there is no evidence of SRF-derived genes in land plants.
Chromatin organization and histone modifications play essential roles in regulating gene expression during development. DEK is a conserved chromatin-associated protein implicated in DNA topology and transcriptional regulation, yet its in vivo function in plants has remained elusive. To uncover DEK function, we used Arabidopsis thaliana dek mutants and performed genome-wide analyses of histone modifications. Genetic and physical interactions with Polycomb Repressive Complex 2 (PRC2)-associated proteins were examined, and transcriptome comparisons were conducted among single and multiple mutants. Loss of DEK function led to a genome-wide increase in the PRC2-mediated histone modification H3K27me3. DEK genetically and physically interacts with the PRC2-associated protein LIKE HETEROCHROMATIN PROTEIN 1 (LHP1), and DEK deficiency partially restores H3K27me3 levels in the lhp1 mutant. The dek multiple mutant exhibited enhanced H3K27me3 at PRC2 target genes and ectopic accumulation at pericentromeric interstitial telomeric repeats - similar to histone H1 mutants - suggesting altered chromatin accessibility. Combined dek and lhp1 mutations intensified developmental defects and disrupted expression of many PRC2 target genes, including MADS-box transcription factors. Transcriptome analyses revealed that DEK and chromatin remodeler alpha thalassemia/mental retardation syndrome X-linked chromatin remodeler have opposing effects on gene expression. Our findings uncover DEK as a novel regulator of H3K27me3 homeostasis and chromatin structure, critical for coordinated plant development.
Proteolysis is a universal process, as proteases play a pivotal role in modulating numerous signaling pathways. Proteases control the fate and function of their target proteins by hydrolyzing peptide bonds within these proteins. Understanding the temporal and spatial dynamics of proteolytic events, including the proteases that execute them, is crucial for elucidating their particular roles across diverse biological processes. In this study, we developed and characterized a set of genetically encoded Förster resonance energy transfer (FRET)-based reporters for the detection of various proteolytic activities in plants. Our sensors reliably reported the activity of specific proteases, exhibiting a performance comparable to previously established detection systems. In addition, we engineered variants capable of detecting the spatial dynamics of metacaspase-triggered proteolysis after wounding and during programmed cell death in roots. We demonstrated the feasibility of these FRET-based sensors for detecting various activities in vivo with high spatiotemporal resolution. The implementation of these tools in plant research opens opportunities to explore proteolytic mechanisms with enhanced precision. Overall, these biosensors constitute a versatile toolbox for probing protease function within its native cellular context, paving the way for deeper insights into plant biology and signaling.
Autophagy is a conserved intracellular catabolic process, critical for plant stress tolerance. Upon their delivery in the vacuole, how autophagic bodies containing cargo are hydrolyzed to warrant autophagy degradation remains unclear in multicellular organisms. Here, we found that two Arabidopsis phospholipases, LCAT4 and LCAT3, traffic to the vacuolar lumen and converge on autophagic bodies through fundamentally different routes. While LCAT4 directly binds ATG8 and uses autophagy as a transport system to reach the vacuole prepackaged within autophagosomes, LCAT3 traffics to the lytic compartment independently of autophagosome formation. Knocking out both genes causes an accumulation of autophagic bodies accompanied with a reduction in autophagy degradation. In vivo reconstitution demonstrated that LCAT3 can hydrolyse the membrane of autophagic bodies, enabling the activity of LCAT4 to enhance this process. Together, this work sheds light on the vacuolar stages of autophagy, showing that plants have evolved a multi-component pathway for the efficient disruption of autophagosomal membranes as a critical step for the completion of the autophagy pathway.